Treasures that scientists keep on ice

Posts Tagged ‘anatomy’

Sorry about the title. It’s the best I could do. In case you missed it on our Anatomy to You blog, we unleashed a hefty database of CT (and some MRI) scans of our frozen crocodile cadavers last week, for free public usage. In total, it’s about 34 individuals from 5 species, in 53 databases constituting around 26,000 individual DICOM file format slices of data. This page has a table of what the data/specimens are. I am writing this post to share some more images and ensure that word gets out. We’re thrilled to be able to finally release this first dataset. We have plans to let loose a LOT more such data in the future, for various organisms that we study.

Stomach-Churning Rating: 2/10- be glad that these data don’t come with an olfactory component, especially the five rotten, maggot-ridden Morelet’s croc specimens, which are among the stinkiest things I’ve dealt with.

Crocodiles are no strangers to this blog, of course, as these past links testify. Indeed, most of the crocodile images I’ve blogged with come from specimens that are in this scan dataset. We even released a “celebrity crocodile, “WCROC” or FNC7 in our dataset, which is the 3.7m long Nile croc from “Inside Nature’s Giants”. It broke our CT scanner back in ~2009 but we got the data, except for the torso, and we also got some MRI scans from it, so we’re chuffed.

Above: The only spectacled caiman (Caiman crocodilus); and indeed the only alligatorid; in our dataset. To watch for: stomach contents/gastroliths, and all the damn osteoderms that I did/didn’t segment in this quickly processed file. This specimen had its limbs dissected for one of our studies, so only the right limbs are visible.

Sharing these kind of huge datasets isn’t so easy. Not only do few websites host them cheaply, and with reasonable file size limits, and limited headaches for what info you have to provide, and with some confidence that the websites/databases will still exist in 5-20 years, but also we were hesitant to release the dataset until we felt that it was nicely curated. Researchers can now visit my lab and study the skeletons (or in some cases, the still-frozen specimens) matched up with the scan data, and known body masses or other metadata. We’re not a museum with dedicated curatorial staff, so that was not trivial to reliably organize, and I still worry that somewhere in the dataset we mis-identified a specimen or something. But we’ve done our best, and I’m happy with that for now.

Above: rotating Osteolaemus tetraspis (specimen FDC2 from our database), which was obviously dissected a bit postmortem before we could scan it, but still shows some cool features like the extensive bony armour and the cute little doglike (to me, anyway) skull. I worked with these animals (live) a bit >10 years ago and came to love them. Compared to some other crocodiles we worked with, they had a pleasant demeanour. Like this guy:

Osteolaemus (resting) set up with motion capture markers for a yet-to-be-published gait study that we did in 2005 (ugh!). It wasn’t harmed by this.

Anyway, as a person who likes to maintain quality in the science we do, I also was hesitant to “just” release the DICOM file data rather than beautiful segmented 3D skeletal (or other tissue) geometry that is ready for 3D printing or animation or other uses, or interactive online tools like Sketchfab. Other labs (e.g. Witmerlab) do these kind of things better than we do and they inspire us to raise our game in the future, but I am sure that we will be forgiven for releasing big datasets without gorgeous visuals and more practical, processed files — this time. 🙂 We agree with many other scientists that sharing data is part of modern, responsible science– and it can be fun, too! Oddly enough, in this case we hadn’t used the CT/MRI data much for our own studies; most of the scans were never fully digitized. We just scan everything we get and figured it was time to share these scans.

Enjoy. If you do something cool with the data that we’ve made accessible, please let us know so we can spread the joy!

And if you’re a researcher headed to ICVM next week, I hope to see you there!

Who needs “Ice Road Truckers” when you have the “John’s Freezer” team on the road with fossils, amphibians, felids and 3D phenotype fun? No one, that’s who. We’re rocking the Cheltenham Science Festival for our first time (as a group), and pulling out all the stops by presenting two events! Here’s the skinny on them, with updates as the week proceeds.

Stomach-Churning Rating: 2/10 for now (just bones), but it could change once the cheetah dissection is under way… 8/10 bloody cheetah bits but only at the end (updated)

Right now, Lauren Sumner-Rooney (of “Anatomy To You” and other fame) is on-site with a rotating team of others from our lab, in the “Free Activity Tents” area of the Imperial Gardens/Square, inside a marquee where we’ll be showing off our NERC-funded tetrapod research all week. This “First Steps” event features not only our past and present work with Jenny Clack, Stephanie Pierce, Julia Molnar and others on Ichthyostega & its “fishapod” mates, but also our “scampering salamanders” research in Spain, Germany and England. I’ve blogged a lot about all that, and won’t repeat it here, but you can see a 3D-printed Ichthyostega skeleton, view the skeleton in a virtual reality 3D environment, see related specimens and engage in kid-friendly activities, and talk to our team about this and other related research.

Ichthyostega 3D printed backbone is born!

The central themes of that event are how bone structure relates to function and how we can use such information, along with experimental measurements and computer models of real salamanders, to reconstruct how extinct animals might have behaved as well as how swimming animals became walking ones. How did fins transform into limbs and what did that mean for how vertebrates made the evolutionary transition onto land? If you know my team’s work, that encapsulates our general approach to many other problems in evolutionary biomechanics (e.g. how did avian bipedalism evolve?). Added benefits are that you too can explore this theme in a hands-on way, and you can talk with us about it in person. That continues all week (i.e. until Saturday evening); I’ll be around from Thursday afternoon onwards, too. Kids of all ages are welcome!

Ichthyostega 3D print taking shape!

Then, on Saturday for our second free event we join forces with Ben Garrod (master of primate evolution, the secrets of bones, and “Attenborough and the Giant Dinosaur”) and RVC’s forensic pathologist Alexander Stoll as well as Sophie Regnault (“sesamoid street” PhD student w/me). As the “Large Animal Dissection” title hints, it’s not the right kind of gig to bring small kids to. There will be blood and stuff— we’ll be dissecting a cheetah together from 10am-4pm. This will involve walking through all the major organ systems, giving evolutionary anecdotes, and plenty more, with an aim to understand how the magnificent adaptations of cheetahs evolved—but also to investigate what problem(s) this animal faced that led to its sad demise. By the day’s end, there will just be a skeleton left. Get a front row seat early for this event, which serendipitously ties into “Team Cat”’s Leverhulme Trust-funded research project (we wanted a big animal and it just happened to be a cheetah; I had hoped for a giant croc or a shark or something but can’t complain!).

Ichthyostega 3D print is ready!

If you miss these events, please do cry bitter tears of regret. But don’t despair, there will be another “big cat dissection” in the London area in ~November (watch here for details), and plenty more fossil tetrapod stuff to come, plus a LOT more dinosaurs on the horizon!

Guess the bones! (photo by Zoe Self)

And please come back to this blog post for more pics and stories as the week carries on… For hashtag afficionados, you can follow the fun on Twitter etc. at #firststepsCSF16. What a world we live in!

Update 3: Cheetah meat & greet
Ben, Alex, Sophie and I tackled the cheetah dissection today and it went GREAT! Much better than I’d optimistically expected. Rain didn’t scare the crowds off and neither did the gore, which there was some of (gelatinous spinal cords, lumpy tumors and at least one flying tiny bit of cheetah flesh that landed on a good-natured audience member!). Photos will tell the tale:

Peek-a-boo!

Sophie and Alex help us get set up in our tent.

Our initial rough schedule- although we ended up improvising more after lunch.

Dissectors assemble!

The beast revealed. It was skinned by the museum that loaned it to us.

Alex showing his talent: removing the viscera in one piece from end to end, starting with the tongue.

Impressive finding of a surgical fixture (plate and wires) on the tibia, which had been used to hold the shattered bone back together long enough for it to heal. Added to the kidney disease and liver-spleen-lung cancer, this cheetah was in the sorriest shape of any cadaver I’ve seen yet.

Dr Adam Rutherford, an eye expert, did a nice dissection of the cheetah’s eye, here showing the tapetum lucidum (reflective membrane), which shows up as light blue colour. Its small size befits the not-very-nocturnal habits of cheetahs.

The lens of the cheetah’s eye. Now cloudy because of dehydration and crystalization, but still fascinating to see.

Happy Darwin Day from the frozen tundra sunny but muddy, frosty lands of England! I bring you limb muscles as peace offerings on this auspicious day. Lots of limb muscles. And a new theme for future blog posts to follow up on: starting off my “Better Know A Muscle” (nod to Stephen Colbert; alternative link) series. My BKAM series intends to walk through the evolutionary history of the coolest (skeletal/striated) muscles. Chuck Darwin would not enjoy the inevitable blood in this photo-tour, but hopefully he’d like the evolution. Off we go, in search of better knowledge via an evolutionary perspective!

There is, inarguably, no cooler muscle than M. caudofemoralis longus, or CFL for short. It includes the largest limb muscles of any land animal, and it’s a strange muscle that confused anatomists for many years– was it a muscle of the body (an axial or “extrinsic” limb muscle, directly related to the segmented vertebral column) or of the limbs (an “abaxial” muscle, developing with the other limb muscles from specific regions of the paraxial mesoderm/myotome, not branching off from the axial muscles)? Developmental biologists and anatomists answered that conclusively over the past century: the CFL is a limb muscle, not some muscle that lost its way from the vertebral column and ended up stranded on the hindlimb.

The CFL is also a muscle that we know a fair amount about in terms of its fossil record and function, as you may know if you’re a dinosaur fan, and as I will quickly review later. We know enough about it that we can even dare to speculate if organisms on other planets would have it. Well, sort of…

Stomach-Churning Rating: 8/10. Lots of meaty, bloody, gooey goodness, on and on, for numerous species. This is an anatomy post for those with an appetite for raw morphology.

Let’s start from a strong (and non-gooey) vantage point, to which we shall return. The CFL in crocodiles and most other groups is (and long was) a large muscle extending from much of the front half or so of the tail to the back of the femur (thigh bone), as shown here:

As the drawing shows, the CFL has a friend: the CFB. The CFB is a shorter, stumpier version of the CFL restricted to the tail’s base, near the hip. The “B” in its name means “brevis”, or runty. It gets much less respect than its friend the CFL. Pity the poor CFB.

But look closer at the CFL in the drawing above and you’ll see a thin blue tendon extending past the knee to the outer side of the lower leg. This is the famed(?) “tendon of Sutton“, or secondary tendon of the CFL. So the CFL has two insertions, one on the femur and one (indirectly) onto the shank. More about that later.

Together, we can talk about these two muscles (CFL and CFB) as the caudofemoralis (CF) group, and the name is nice because it describes how they run from the tail (“caudo”) to the femur (“femoralis”). Mammal anatomists were late to this party and gave mammal muscles stupidly unhelpful names like “gluteus” or “vastus” or “babalooey”. Thanks.

But enough abstract drawings, even if they rock, and enough nomenclature. Here is the whopping big CFL muscle of a real crocodile:

Huge Nile crocodile, but a relatively small CFL.

Bigger crocs have smaller legs and thus smaller leg muscles, relatively speaking. CFL at the top, curving to the left.

The giant Nile croc’s CFL muscle removed for measurements. 2.35 kg of muscle! Not shabby for a 278 kg animal.

However, maybe crocodile and other archosaur CFL muscles are not “average” for leggy vertebrates? We can’t tell unless we take an evolutionary tack to the question.

Where did the CFL come from, you may ask? Ahh, that is shrouded in the fin-limb transition‘s mysteries. Living amphibians such as salamanders have at least one CF muscle, so a clear predecessor to the CFL (and maybe CFB) was present before reptiles scampered onto the scene.

But going further back through the CF muscles’ history, into lobe-finned fish, becomes very hard because those fish (today) have so few fin muscles that, in our distant fishy ancestors, would have given rise eventually to the CF and other muscle groups. With many land animals having 30+ hindlimb muscles, and fish having 2-8 or so, there obviously was an increase in the number of muscles as limbs evolved from fins. And because a limb has to do lots of difficult three-dimensional things on land while coping with gravity, more muscles to enable that complex control surely were needed.

OK, so there were CF muscles early in tetrapod history, presumably, anchored on that big, round fleshy tail that they evolved from their thin, finned fishy one — but what happened next? Lizards give us some clues, and their CFL muscles aren’t all that different from crocodiles, so the CFL’s massive size and secondary “tendon of Sutton” seems to be a reptile thing, at least.

Courtesy of Emma Schachner, a large varanid lizard’s very freshly preserved CFL and other hindlimb muscles.

Courtesy of Emma Schachner, zoomed in on the tendons and insertions of the CFL muscle and others. Beautiful anatomy there!

Looking up at the belly of a basilisk lizard and its dissected right leg, with the end of the CFL labelled. It’s not ideally dissected here, but it is present.

An unspecified iguanid(?) lizard, probably a juvenile Iguana iguana, dissected to reveal its CFL muscle near its attachment to the femur. The muscle would extend further, about halfway down the tail, though.

Let’s return to crocodiles, for one because they are so flippin’ cool, and for another because they give a segue into archosaurs, especially dinosaurs, and thence birds:

A moderate-sized (45kg) Nile crocodile with its CFL muscle proudly displayed. Note the healthy sheath of fat (cut here) around the CFL.

American alligator’s CFL dominates the photo [by Vivian Allen].

Black caiman, Melanosuchus, showing off its CFL muscle (pink “steak” in the middle of the tail near the leg), underneath all that dark armour and fatty superficial musculature.

A closer look at the black caiman’s thigh and CFL muscle.

Like I hinted above, crocodiles (and the anatomy of the CFL they share with lizards and some other tetrapods) open a window into the evolution of unusual tail-to-thigh muscles and locomotor behaviours in tetrapod vertebrates.

Thanks in large part to Steve Gatesy’s groundbreaking work in the 1990s on the CFL muscle, we understand now how it works in living reptiles like crocodiles. It mainly serves to retract the femur (extend the hip joint), drawing the leg backwards. This also helps support the weight of the animal while the foot is on the ground, and power the animal forwards. So we call the CFL a “stance phase muscle”, referring to how it mainly plays a role during ground contact and resisting gravity, rather than swinging the leg forwards (protracting the limb; i.e. as a “swing phase muscle”).

The “tendon of Sutton” probably helps to begin retracting the shank once the thigh has moved forward enough, facilitating the switch from stance to swing phase, but someone really needs to study that question more someday.

And thanks again to that same body of work by Gatesy (and some others too), we also understand how the CFL’s anatomy relates to the underlying anatomy of the skeleton. There is a large space for the CFL to originate from on the bottom of the tail vertebrae, and a honking big crest (the fourth trochanter) on the femur in most reptiles that serves as the major attachment point, from which the thin “tendon of Sutton” extends down past the knee.

Femur bones (left side; rear view) from an adult ostrich (left) and Nile crocodile (right). Appropriate scale bar is appropriate. The fourth trochanter for the CFL is visible in the crocodile almost midway down the femur. Little is left of it in the ostrich but there is a bumpy little muscle scar in almost the same region as the fourth trochanter, and this is where the same muscle (often called the CFC; but it is basically just a small CFL) attaches.

That relationship of the CFL’s muscular anatomy and the underlying skeleton’s anatomy helps us a lot! Now we can begin to look at extinct relatives of crocodiles; members of the archosaur group that includes dinosaurs (which today we consider to include birds, too), and things get even more interesting! The “tendon of Sutton”, hinted at by a “pendant” part of the fourth trochanter that points down toward the knee, seems to go away multiple times within dinosaurs. Bye bye! Then plenty more happens:

A large duckbill dinosaur’s left leg, with a red line drawn in showing roughly where the CFL would be running, to end up at the fourth trochanter. Many Mesozoic dinosaurs have skeletal anatomy that indicates a similar CFL muscle.

We can even go so far as to reconstruct the 3D anatomy of the CFL in a dinosaur such as T. rex (“Sue” specimen here; from Julia Molnar’s awesome illustration as part of our 2011 paper), with a fair degree of confidence. >180kg steak, anyone?

As we approach birds along the dinosaur lineage, the tail gets smaller and so does the fourth trochanter and thus so must the CFL muscle, until we’re left with just a little flap of muscle, at best. In concert, the hindlimbs get more crouched, the forelimbs get larger, flight evolves and voila! An explosion of modern bird species!

Left femur of an ostrich in side view (hip is toward the right side) showing many muscles that attach around the knee (on the left), then the thin strap of CF muscle (barely visible; 2nd from the right) clinging near the midshaft of the femur.

Another adult ostrich’s CF muscle complex, removed for study. Not enough ostrich myology for you yet? Plenty more in this old post!Or this one!Or this one… hey maybe I need to write less about ostriches? The CF muscle complex looks beefy but it’s no bigger than any other of the main hindlimb muscles, unlike the CFL in a crocodile or lizard, which puts everything else to shame!

STILL not enough ostrich for you yet? Take a tour of the major hindlimb muscles in this video:

And check out the limited mobility of the hip joint/femur here. No need for much femur motion when you’re not using your hip muscles as much to drive you forwards:

But I must move on… to the remainder of avian diversity! In just a few photos… Although the CF muscles are lost in numerous bird species, they tend to hang around and just remain a long, thin, unprepossessing muscle:

Chicken’s right leg in side view. CFC muscle (equivalent of CFL; the ancestral CFB is confusingly called the CFP in birds, as it entirely resides on the pelvis) outlined and labelled.

A jay (species? I forget) dissected to show some of the major leg muscles, including the CFL-equivalent muscle; again, smallish. [Photo by Vivian Allen]

Finally, what’s up with mammals‘ tail-to-thigh CF-y muscles? Not much. Again, as in birds: smaller tail and/or femur, smaller CF muscles. Mammals instead depend more on their hamstring and gluteal muscles to support and propel themselves forward.

But many mammals do still have something that is either called the M. caudofemoralis or is likely the same thing, albeit almost always fairly modest in size. This evolutionary reduction of the CF muscle along the mammal (synapsid) lineage hasn’t gotten nearly as much attention as that given to the dinosaur/bird lineage’s CFL. Somebody should give it a thoroughly modern phylogenetic what-for! Science the shit outta that caudofemoralis…

Cats have a pretty large CF muscle in general, and this jaguar is no exception! But mammals still tend to have fairly wimpy tails and thus CF muscles, or they even lose them (e.g. us?). [photo by Andrew Cuff, I think]

In summary, here’s what happened (click to embeefen):

Better Know A Muscle: the evolution of M. caudofemoralis (longus).

I hope you enjoyed the first BKAM episode!
I am willing to hear requests for future ones… M. pectoralis (major/profundus) is a serious contender.

P.S. It was Freezermas this week! I forgot to mention that. But this post counts as my Freezermas post for 2016; it’s all I can manage. Old Freezermas posts are here.

Tonight is the debut of the ballyhooed BBC1 programme “Attenborough and the Giant Dinosaur“, featuring Sir David and The Titanosaur-With-No-Name, at 6:30pm. Furthermore, this week I presided over a showing of “T. rex Autopsy” to our RVC undergrad Zoological Society, with a very enjoyable Q&A afterwards. So it seemed timely for me to do a post about a theme that links these two documentaries that I helped with, my own research, and science communication and palaeontological research more generally. But first let’s get this out of the way:

It was great. I could gush more but that’s not what this post is about.

For the titanosaur documentary with Sir David, and the incomparable Ben Garrod as well, we had an old elephant “friend” of mine (subject of many biomechanics studies we’d done) walk across a pressure pad to demonstrate how the elephant locomotion works and some of the basic similarities with how a giant sauropod dinosaur might walk:

A broader feature of that documentary, which elephants are linked into, is how we can use the skeleton to reconstruct some general aspects of the soft tissue anatomy, and thereby the physiology or even behaviour, of a giant titanosaur. Which brings me to this post’s subject: We dig up dinosaurs all the time, but what about digging into dinosaurs and using what’s preserved to reconstruct what isn’t?

Some of the “G-suit” compressive stocking that the fascia wrapped around elephant, and other large mammals, creates, and the underlying, interwoven muscles and tendons (hindlimb of a young Asian elephant that sadly died in captivity). Did some larger dinosaurs have something like this? I’d expect so.

Another view, more superficially, of that G-suit/stocking under the thick, tight skin of an elephant’s leg. You’ll hear more about this in the Attenborough show…

Once the documentary airs, I may be able to share some more images from it showing what they did for the titanosaur, but this BBC photo gives a good idea. Here, blood vessels and other tissues surrounding the skeleton. How would a titanosaur pump blood around its body? A good question.

I wanted to focus more on the end product; the imagery or even physical object; and how it conveys what we think we know about dinosaur anatomy. I’ll do that via examples, using photos of dinosaur anatomy that I’ve collected over the years from museums or other media. There won’t be any profound points or long musings; it’s mainly a photo blog:

What your (inner?) child most needs is a dinosaur to dissect yourself! Why not a T. rex toy like this?

For the price (~$30 in USA), the 4D Vision dinosaurs deliver a pretty good bargain, and the anatomy is satisfactory. I’ve been collecting this series. I could quibble, but hey: it’s a dinosaur you get to build/dissect yourself, and with many major organs in reasonable positions! Not so easy to put/keep together, but it’s fine. I would not pay a ton for it, though.

Closeup of the leg muscles- hey, not bad! Pretty much the right muscles in the right places more or less, and plausible proportions. No air sacs in the torso, but again, this is mid-1990’s science shown. BUT…

I was happy with this poster until I got it home from the western-USA museum I bought it at and read this final bit of text… Oh, America! You silly place. (And unfortunately, these dinosaurs are not from the very end of the Cretaceous anyway, so “global catastrophe” is not implicated.)

Looking down onto the opened torso of the Barcelona ornithomimid. Strikingly realistic! Breastbone, heart, liver, intestines; not unreasonable positions and sizes. No feathers, but again hey– this was made in the earlier days.

Albertosaurus from the Drexel Academy of Sciences. I forget where I got this pic but I like the display– this is an impressive full-scale physical model. The transition from skeleton-only on the left to skinned in the middle to fully-fleshed and popping out atcha on the right is clever.

?T. rex? leg, photo that I took ages ago as a PhD student, if memory serves. Can anyone remind me where this was? California Academy of Sciences? I am embarrassed that I cannot recall. I remember geeking out about it. It has a toy-ish look, but I reckon they had to build this to withstand kids touching it.

Perhaps the best dino-dissection example I’ve seen in a museum: the AMNH’s sauropod Mamenchisaurus with internal organs and their functions projected onto it, in the “World’s Largest Dinosaurs” exhibit. Bravo! I stood and watched it for quite a while.

This is far from comprehensive– just several kinds of imagery that I mostly like. There’s the tension between showing too much realism, which science simply can’t back up, and being too cartoonish, losing the viewer’s immersion in the time-travelling fantasy. I do, however, like other kinds of more abstract, schematic depictions of dinosaur anatomy that simplify the details to focus on the basics of what organs should have been where and how they may have worked, such as this depiction from T. rex Autopsy, which also took the other extreme favouring ultra-realism (but with physical models, not so much with the CGI):

Air flow through a T. rex: simplified but clear. CGI used to explain, not abused. The real air sac anatomy would be too complex to show. You may see something similar with the titanosaur show.

That’s enough for now. I’ve stuck with relatively recent examples; of course in my particular field I also think back to Romer’s wonderful 1920’s drawings, which I covered in this post.

So, blog readers, help me out here: what examples of dinosaur internal, squishy anatomy from museums, documentaries or other not-entirely-done-by-nitpicky-scientists venues do you like, or not like so much? What works for you, or at least is memorable in some way?

Greetings Freezerinos, and Happy New Year! I have been quiet on this blog for health and other reasons but those will pass and there will be new posts in 2016. However, behind the scenes there have been super-cool things afoot. I am very happy to bring one of them to you now:

We have just debuted our new social media “presence” (for lack of a better word) that is a sister blog to this one. It is called Anatomy To You (http://anatomytoyou.com/), as its intent is to bring a wide array of science about animal anatomy to “you”, the general public. This John’s Freezer blog will continue with it’s style of rambling longer posts targeted at a fairly geeky scientifically literate audience and focusing on my team’s research and my own disparate thoughts about science and related issues. Anatomy To You will bring you shorter posts, even just images, completely focused on celebrating the structure of organisms, and not just presenting my team’s research but also a wide array of anatomical science from around the globe. It will also be much more regular and frequent in its posts. We’ll welcome guest posts and I encourage you to get in touch with us if you want to jump on the bandwagon early, or have us feature your research for you!

Anatomy To You will continue to evolve over this coming year, so please stay with us and give us feedback; join in the morphological conversations with us. I am SUPER excited to see where this goes– it is an experiment that has a lot of potential, we think.

Seeking adaptations for running and swimming in the vertebral columns of ancient crocs

A guest post by Dr. Julia Molnar, Howard University, USA (this comes from Julia’s PhD research at RVC with John & colleagues)

Recently, John and I with colleagues Stephanie Pierce, Bhart-Anjan Bhullar, and Alan Turner described morphological and functional changes in the vertebral column with increasing aquatic adaptation in crocodylomorphs (Royal Society Open Science, doi 10.1098/rsos.150439). Our results shed light upon key aspects of the evolutionary history of these under-appreciated archosaurs.

Stomach-Churning Rating: 5/10; a juicy croc torso in one small photo but that’s all.

Phylogenetic relationships of the three crocodylomorph groups in the study and our functional hypotheses about their vertebrae. * Image credits: Hesperosuchus by Smokeybjb, Suchodus by Dmitry Bogdanov (vectorized by T. Michael Keesey) http://creativecommons.org/licenses/by-sa/3.0

As fascinating as modern crocodiles might be, in many ways they are overshadowed by their extinct, Mesozoic cousins and ancestors. The Triassic, Jurassic, and early Cretaceous periods saw the small, fast, hyper-carnivorous “sphenosuchians,” the giant, flippered marine thalattosuchians, and various oddballs like the duck-billed Anatosuchus and the aptly named Armadillosuchus. As palaeontologists/biomechanists, we looked at this wide variety of ecological specializations in those species, the Crocodylomorpha, and wanted to know, how did they do it?

Of course, we weren’t the first scientists to wonder about the locomotion of crocodylomorphs, but we did have some new tools in our toolbox; specifically, a couple of micro-CT scanners and some sophisticated imaging software. We took CT and micro-CT scans of five fossil crocodylomorphs: two presumably terrestrial early crocodylomorphs (Terrestrisuchus and Protosuchus), three aquatic thalattosuchians (Pelagosaurus, Steneosaurus, and Metriorhynchus) and a semi-aquatic modern crocodile (Crocodylus niloticus). Since we’re still stuck on vertebrae (see, e.g., here; and also here), we digitally separated out the vertebrae to make 3D models of individual joints and took measurements from each vertebra. Finally, we manipulated the virtual joint models to find out how far they could move before the bones bumped into each other or the joints came apart (osteological range of motion, or RoM).

Our methods: get fossil (NHMUK), scan fossil, make virtual fossil and play with it.

Above: Video of a single virtual inter-vertebral joint from the trunk of Pelagosaurus typus (NHMUK) showing maximum osteological range of motion in the lateral direction (video). Note the very un-modern-croc-like flat surfaces of the vertebral bodies! (modern crocs have a ball-and-socket spinal joint with the socket on the front end)

While this was a lot of fun, what we really wanted to find out was whether, as crocodylomorphs became specialized for different types of locomotion, the shapes of their vertebrae changed similarly to those of mammalian lineages. For example, many terrestrial mammals have a lumbar region that is very flexible dorsoventrally to allow up-and-down movements during bounding and galloping. Did fast-running crocodylomorphs have similar dorsoventral flexibility? And did fast-swimming aquatic crocodylomorphs evolve a stiffer vertebral column like that of whales and dolphins?

Above: Video of how we modelled and took measurements from the early crocodylomorph Terrestrisuchus gracilis (NHMUK).

Our first results were puzzling. The Nile croc had greater RoM in side-to-side motions, which makes sense because crocodiles mostly use more sprawling postures and are semi-aquatic, using quite a bit of side-to-side motions in life. The part that didn’t make sense was that we found pretty much the same thing in all of the fossil crocodylomorphs, including the presumably very terrestrial Terrestrisuchus and Protosuchus. With their long limbs and hinge-like joints, these two are unlikely to have been sprawlers or swimmers!

So we started looking for other parts of the croc that might affect RoM. The obvious candidate was osteoderms, the bony scales that cover the back. We went back to John’s Freezer and got out a nice frozen crocodile to measure the stiffness of its trunk and found that, sure enough, it was a lot stiffer and less mobile without the osteoderms. If the fairly flexible arrangement of osteoderms in crocodiles had this effect on stiffness, it seemed likely that (as previous authors have suggested; Eberhard Frey and Steve Salisbury being foremost amongst them) the rigid, interlocking osteoderms running from head to tail in early crocodylomorphs would really have put the brakes on their ability to move their trunk in certain ways.

Testing the stiffness of (Nile) crocodile trunks to learn the effects of osteoderms, skin, muscles, and ribs. We hung metric weights from the middle of the trunk and measured how much it flexed (Ɵ), then removed bits and repeated. Click to em-croccen.

Another cool thing we found was new evidence of convergent evolution to aquatic lifestyles in the spines of thalattosuchians. The more basal thalattosuchians, thought to have been near-shore predators, had stiffness and RoM patterns similar to Crocodylus. But Metriorhynchus, which probably was very good at chasing down fast fish in the open ocean, seems to have had greater stiffness. (The stiffness estimates come from morphometrics and are based on modern crocodiles; see here again, or just read the paper already!) A stiff vertebral column can be useful for a swimmer because it increases the body’s natural frequency of oscillation, and faster oscillation means faster swimming (think tuna, not eel). The same thing seems to have happened in other secondarily aquatic vertebrate lineages such as whales, ichthyosaurs, and mosasaurs.

So, our results were a mixed bag of adaptations particular to crocs and ones that seem like general vertebrate swimming specializations. Crocodylomorphs are important because they are the only group of large vertebrates other than mammals that has secondarily aquatic members and has living members with a reasonably similar body plan, allowing us to test hypotheses in ways that would arguably be impossible for, say, non-avian dinosaurs and birds. The take-home message: crocodylomorphs A) are awesome, and B) can teach us a lot about how vertebrates adapt to different modes of life.

I was recently featured on Daily Planet, a great Canadian science show on TV that lamentably is not broadcast more globally. It is always high quality science communication, aided by the superb hosts Ziya Tong and Dan Riskin (and a talented crew!). What were we doing? Dissecting an elephant’s foot, of course!

Stomach-Churning Rating: 9/10; no-holds-barred dismantling of elephant feet, from the video onwards, and this post is heavy on moist, goopy photos afterwards, with some nasty pathologies. Not nice at all. I’ll give you a chance to turn around while contemplating the cart that we use to carry elephant feet around campus (each foot is 20-30kg; up to 70lbs; so we need the help!), before the video.

Here is a snippet of the full segment from Daily Planet:

And here is more of some of my recent dissections. I’ll walk you through two dissections, via photos. This goes back to the roots of this blog: unflinching, gritty examinations of real anatomy! Of course, no elephants were harmed for this work. They died at EU zoos/parks and were sent to me for postmortem examination and research, so we hope that this benefits the future care of elephants. We’re currently finishing up a grand overview paper that describes all of the odd pathologies we’ve observed in elephant feet, for the benefit of zoo keepers and vets who are trying to detect, diagnose and monitor any foot problems.

There are two feet in this post, both front right feet (manus is the technical term; singular and plural). The first one is the messier (unhealthy and bloodier, less fresh and clean) one, from the show/video. It is an Asian elephant (Elephas maximus). I kick off with photos I took after the filming, so the foot is already deconstructed:

Skinned foot, oblique front/inside view. The wrist is on the right side of the photo; the toes on the left.

Sole (“slipper”), with a hole on the fourth toe showing where the abscess is that let infection in/pus drain out. The slipper here is upside-down.

Top-down view of the sole of the foot, once the slipper is removed; flipped over and rotated 90 degrees clockwise from the above photo. Some of the fat pad of the foot is on the right side of the image; it’s very hard to separate from the keratinous sole of the foot.

Looking down into the fourth toe’s (ring finger) abscess on the other side of the above view.

Looking down into the second toe (index finger), same view as above. Some redness and greyness where this toe had some of its own pathological issues like infection and a smaller abscess.

Looking up from the slipper (removed) at the fat pad and toes of the foot, where they interface with the sole/slipper. The fat pad is toward the bottom and left side; the five toes are on the upper/right side (knobby subcircular regions on the perimeter of the foot). The very bad infection on the fourth toe is visible on the bottom right.

The sproingy fat pad is worth a video!

And one good wiggle deserves another!

A view down onto the wrist joint. The carpal (wrist) bones are visible at the bottom of the image, whereas the flexor (palmar) tendons and muscles on the back of the “hand” are at the top. There is a LOT of musculotendinous tissue on the back side of an elephant’s foot. As you will see in my dissection of the second foot, further below!

Looking down onto the medial (inner/”thumb”) border of the foot, where I’ve exposed the prepollex, or false “sixth finger”, by removing the first metacarpal (knuckle) bone.

I’ve removed the prepollex from the foot. The white oval structure (bottom right) is the top of the conical prepollex, where it connected to the rest of the foot. White is cartilage, whereas the red “islands” are blood vessels that have invaded the cartilage and are starting to turn it into patches of bone. So this prepollex is at a very early stage of bone formation, still almost entirely cartilaginous, whereas some older elephants have the prepollex largely formed of bone. The fleshy pink tissue adhering to the surface of the prepollex here is a remnant of “abductor” muscle that connects it to the thumb and thus could allow some active control of the prepollex’s mobility.

Well, that was one very pathological elephant’s foot; one of the worst I have ever seen. Every foot I dissect is different and tells me a unique story about that animal’s development, history and health. This one told a very sad tale. What does a somewhat normal elephant’s foot look like? I thawed one out for comparison, and to thin out my overstuffed freezer stock. This one starts off from an intact (if severed) foot so you can witness the stages of dissection:

Whole foot. African elephant (Loxodonta africana). You may spot in later photos that the second and fourth toes’ nails are cracked longitudinally. This happens sometimes in elephants without any obvious health problems such as infection, but if it lasts long enough and conditions are bad enough (e.g. unsanitary conditions getting bacteria into the crack; spreading the crack to let them into the foot tissue), it could worsen.

Nice clean sole. No abscesses or other problems. You can faintly see the cracked toenails here.

Gorgeous white cartilage surfaces of the wrist joints. Nice and healthy-looking. A young animal, in this case.

Skinned foot; front view. The thin, broad extensor tendons that would draw the fingers forward in life are visible here as longitudinal lines along the foot’s surface, running to the toes.

Ahh, my favourite thing! I’ve cut around the prepollex and am pointing at it. It’s almost impossible otherwise to see through all the fatty tissue of the fat pad that surrounds it.

Removing the prepollex. It’s tiny and enmeshed in connective tissue; harder to see than in the first elephant (photos above).

There is the prepollex! Maybe 12cm long. A little bit of cartilage (white) visible where it connected to the foot. These “sesamoid bones” vary tremendously in elephants I’ve inspected. I am still getting my head around that, after >10 years of staring at them in >75 feet!

Gap left by removal of the prepollex, on the median border of the foot; thumb region. Imagine having a little extra thumb growing off the base of your thumb and sticking toward your palm. That’s what elephants have.

Here, removing the slipper/sole of the foot, from the back side forwards. Hard work!

The slipper. Compare with the image above (same orientation). Nothing wrong here that I could see.

Front view of the toes, where they connect to the toenails. This specimen was so fresh that they were surprisingly easy to cut through and remove the foot from the sole.

Looking up at the palm. You can see the bulbous fat pad (yellower tissue) bulging out in the centre of the palm, and segments of it extending between each finger, separated by fibrous tracts. I love this anatomy. I can stare at it for hours and still be fascinated after all these years. So complex!

Looking down onto the inside of the toenails, toes 3 and 4. Healthy, relatively intact tissue; no swelling or bleeding or other pathology.

Skinned foot, oblique front/inside view again, as above.

Fat pad removed, looking up through where it was at the palm of the “hands”, where the tendons and ligaments connect to the five toes. Each arc-like structure is a toe; the “thumb” (first toe) is on the upper left.

Elephant’s-eye-view looking down onto the fat pad, where the palm of the foot in the image below would be placed in life (i.e. the limb would be coming down vertically, perpendicular to the plane of the image). The fat pad of the foot is visibly thicker toward the back of the foot (bottom of the image), as you’d expect, because the toes occupy most of the front parts.

Palmar tendons and muscles; the common digital extensor muscle group, which clenches the toes. Not a small muscle, either!

Tendons of the digital flexor muscle exposed.

I removed the digital flexor muscle so the three major tendons can be seen (the two short side branches to the first and fifth toes have been cut off).

Forefoot with flexor tendons removed, revealing the channels that they coursed through.

Closeup of the glistening channels for the flexor tendons. They are lined with lubricative tissue to help the tendons glide through them. And the tendons do need to be able to glide- although elephant feet look very solid from the outside, and are to an extent, but we’ve done studies showing that they do move if you apply even a moderate load to them in a cadaver, and thus would move in life, too.

Let’s finish off with some osteology, shall we? First the unhealthy Asian elephant, then the healthy African elephant; same front right feet, just the bones (from my CT scans):

Ouch, indeed!

Much better. And that’s the end!

Wow, that was an elephantine post! I wanted to take yet another opportunity to share the amazing anatomy of elephant feet with you. You’re all now qualified experts if you made it this far!